The coccidian genus Caryospora

Compiled by
Donald W. Duszynski and Lee Couch
Department of Biology, University of New Mexico
and
Steve J. Upton
Division of Biology, Kansas State University

Supported by NSF-PEET DEB 9521687

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Updated: 24 March 2003
Comments: eimeria@unm.edu or coccidia@ksu.edu


The coccidian genus Caryospora Leger, 1904 is the third largest genus in the Eimeriidae. Oocysts possess a single sporocyst, each sporocyst with 8 sporozoites. The majority of species occur either in raptors or snakes, and the morphologic differences suggest independent lineages. Most species are homoxenous, although some are facultatively homoxenous and can utilize rodents as transport/intermediate hosts. Experimental transmission studies using oocysts collected from colubrids and viperids suggests the former are homoxenous whereas the latter are facultatively heteroxenous.


Class: Aves

  1. Caryospora aquilae Volf, Koudela, and Modry, 2000
    1. Synonym: Caryospora sp. 2 of Volf and Modry, 1998
    2. Host: Aquila chrysaetos (Golden eagle - Falconiformes)
    3. References: Volf and Modry 1998; Volf et al. 2000

  2. Caryospora arcayae Volcan and Medrano, 1984
    1. Synonym: Caryospora sp. of Upton and Sundermann, 1990
    2. Hosts: Buteo magnirostris (Roadside hawk - Falconiformes), Buteo platypterus (Broad-winged hawk - Falconiformes)
    3. References: Upton and Sundermann 1990; Upton et al. 1990; Volcan and Medrano (1984)

  3. Caryospora argentati Schwalbach, 1959
    1. Host: Larus argentatus (Herring gull - Charadriiformes)
    2. Reference: Schwalbach 1959
    3. This coccidian could also be easily placed within the genus Tyzzeria

  4. Caryospora boeri Kluh, 1994
    1. Synonym: Caryospora sp. of Kutzer, Frey, and Kotremba, 1980
    2. Host: Falco tinnunculus (Eurasian or common kestrel - Falconiformes)
    3. References: Kluh 1994; Kutzer et al. 1980

  5. Caryospora bubonis Cawthorn and Stockdale, 1981
    1. Host: Bubo virginianus (Great horned owl - Strigiformes)
    2. References: Cawthorn and Stockdale 1981, 1982; Stockdale and Cawthorn 1981

  6. Caryospora circi Volf, Koudela, and Modry, 2000
    1. Synonym: Caryospora sp. 1 of Volf and Modry, 1998
    2. Host: Circus aeruginosus (Marsh harrier - Falconiformes)
    3. References: Volf and Modry 1998; Volf et al. 2000

  7. Caryospora falconis Wetzel and Enigk, 1937
    1. Hosts: Falco peregrinus (Peregrine falcon), Falco subbuteo (European hobby), Falco tinnunculus (Eurasian or common kestrel) (all Falconiformes)
    2. References: Frey and Kutzer (1982); Kluh 1994; Kutzer et al. (1980); Wetzel and Enigk (1937; 1939); Yakimoff and Matschoulsky (1936)

  8. Caryospora gloriae Pellerdy, 1967
    1. Host: Dives atroviolaceus (Cuban blackbird - Passeriformes)
    2. Reference: Pellerdy 1967
    3. This may be an erroneous description based on improper sporulation of an isosporan

  9. Caryospora henryae (Yakimoff and Matikaschwili, 1932) Yakimoff and Matschulsky, 1936
    1. Synonym: Isospora henryi Yakimoff and Matikaschwili, 1932
    2. Host: Bubo bubo (Northern eagle owl - Strigiformes)
    3. References: Yakimoff and Matikachwili (1932); Yakimoff and Matschoulsky (1936, 1940)

  10. Caryospora jiroveci Cerna, 1976
    1. Host: Erithacus rubecula (European robin - Passeriformes)
    2. References: Cerna 1976; Svobodova 1994

  11. Caryospora kansasensis Upton, Campbell, Weigel, and McKown, 1990
    1. Synonym: Caryospora sp. of Upton and Sundermann, 1990
    2. Host: Buteo swainsoni (Swainson's hawk - Falconiformes)
    3. References: Upton and Sundermann 1990; Upton et al. 1990

  12. Caryospora kutzeri Boer, 1982
    1. Synonym: Caryospora falconis of Schellner and Rodler, 1971
    2. Synonym: Caryospora henryi of Yakimoff and Matschoulsky, 1936, pro parte
    3. Synonym: Caryospora sp. of Cringoli, Quesada, and Papparella, 1991
    4. Hosts: Falco biarmicus (Lanner falcon), Falco cherrug (Saker falcon), Falco jugger (Lagger falcon), Falco mexicanus (Prairie falcon), Falco peregrinus (Peregrine falcon), Falco rusticolus (Gyrfalcon), Falco subbuteo (European hobby), Falco tinnunculus (Eurasian or common kestrel) (all Falconiformes)
    5. References: Boer 1982; Cringoli et al. 1989, 1991; Kluh 1994; Schaffrath-Boer 1983; Schellner and Rodler 1971; Volf and Modry 1998; Yakimoff and Matschoulsky 1936

  13. Caryospora lindsayi Upton, Campbell, Weigel, and McKown, 1990
    1. Host: Buteo jamaicensis (Red-tailed hawk - Falconiformes)
    2. Reference: Upton et al. 1990

  14. Caryospora megafalconis Kluh, 1994
    1. Hosts: Falco cherrug (Saker falcon), Falco rusticolus (Gyrfalcon), Falco tinnunculus (Eurasian or common kestrel) (all Falconiformes)
    2. References: Kluh 1994; Kutzer et al. 1980

  15. Caryospora neofalconis Boer, 1982
    1. Hosts: Falco biarmicus (Lanner falcon), Falco cherrug (Saker falcon), Falco mexicanus (Prairie falcon), Falco peregrinus (Peregrine falcon), Falco subbuteo (European hobby), Falco tinnunculus (Eurasian or common kestrel) (all Falconiformes)
    2. References: Boer 1982; Kluh 1994; Volf and Modry 1998

  16. Caryospora sp. of Svobodova, 1994
    1. Host: Acrocephalus palustris (Marsh warbler - Passeriformes)
    2. References: Svobodova 1990, 1994

  17. Caryospora sp. of Svobodova, 1994
    1. Host: Hippolais sp. (Passeriformes)
    2. Reference: Svobodova 1994

  18. Caryospora sp. of Varghese and Yayabu, 1981
    1. Host: Diphyllodes magnificus (Magnificent bird of paradise - Passeriformes)
    2. References: Varghese and Yayabu 1981
    3. This is probably an isosporan that sporulated abnormally

  19. Caryospora sp. of Wetzel and Enigk, 1939
    1. Synonym: Caryospora falconis of Wetzel and Enigk, 1939
    2. Host: Athene noctua (Little owl - Strigiformes)
    3. References: Wetzel and Enigk 1939

  20. Caryospora sp. of Yakimoff and Matschoulsky, 1936
    1. Synonym: Caryospora henryae of Yakimoff and Matschoulsky, 1936, pro parte (ovoid form)
    2. Host: Milvus migrans (Black kite - Falconiformes)
    3. Reference: Yakimoff and Matschoulsky 1936

  21. Caryospora sp. of Yakimoff and Matschoulsky, 1936
    1. Synonym: Caryospora henryae of Yakimoff and Matschoulsky, 1936, pro parte (round form)
    2. Host: Milvus migrans (Black kite - Falconiformes)
    3. Reference: Yakimoff and Matschoulsky 1936

  22. Caryospora strigis Gottschalk, 1972
    1. Host: Tyto alba (Barn owl - Strigiformes)
    2. References: Gottschalk 1972

  23. Caryospora tremula (Allen, 1933) Hoare, 1934
    1. Synonym: Eumonospora tremula Allen, 1933
    2. Host: Cathartes aura (Turkey vulture - Falconiformes)
    3. References: Allen 1933, 1934; Hoare 1934

  24. Caryospora undata Schwalbach, 1959
    1. Synonym: Caryospora undulata Poelma and Strik, 1966, lapsus
    2. Hosts: Larus argentatus (Herring gull), Lunda cirrhata (Tufted puffin), Uria aalge aalge (Common murre or guillemot) (all Charadriformes)
    3. References: Poelma and Strik 1966; Schwalbach 1959; Upton and Sundermann 1990; Upton et al. 1992
    4. This coccidian could also be easily placed within the genus Tyzzeria

  25. Caryospora uptoni Lindsay and Blagburn, 1986
    1. Host: Buteo jamaicensis (Red-tailed hawk - Falconiformes)
    2. References: Lindsay and Blagburn 1986, 1989; Upton et al. 1990

References for the Caryospora spp. of Aves

ALLEN, E.A. 1933. Eumonospora tremula gen. et sp. nov., a coccidium from the intestine of the turkey buzzard, Cathartes aura septentrionalis Weid. Transactions of the American Microscopical Society 52: 192-194.

ALLEN, E.A. 1934. Eumonospora tremula (Allen, 1933) emended to Caryospora tremula (Allen, 1933) Hoare, 1934. Transactions of the American Microscopical Society 53: 293.

BOER, B. 1982. Untersuchungen uber das Vorkommen von Kokzidien bei Greifvogeln und uber die Entwicklung von zwei Caryospora-Arten der Falken (Caryospora neofalconis n. sp. und Caryospora kutzeri n. sp.). Dissertation, Institute fur Parasitologie der Tierarzlichen Hochschule Hannover, 83 pp.

BURTSCHER, H. 1966. Nieren-Kokzidiose bei Eulen. Wiener Tierarztlichen Monatschrift 53: 654-666.

CAWTHORN, R.J. and STOCKDALE, P.H.G. 1981. Description of Eimeria bubonis sp. n. (Protozoa: Eimeriidae) and Caryospora bubonis sp. n. (Protozoa: Eimeriidae) in the great horned owl, Bubo virginianus (Gmelin), of Saskatchewan. Canadian Journal of Zoology 59: 170-173.

CAWTHORN, R.J. and STOCKDALE, P.H.G. 1982a. The developmental cycle of Caryospora bubonis Cawthorn and Stockdale 1981 (Protozoa: Eimeriidae) in the great horned owl, Bubo virginianus (Gmelin). Canadian Journal of Zoology 60: 152-157.

CERNA, Z. 1973. Interesting coccidians from wild birds. Journal of Protozoology, Proceedings of the Society of Protozoologists 20 (Abstract 136): 535-536.

CERNA, Z. 1974. Two new coccidians from passeriform birds. Folia Parasitologica (Praha) 23: 277-279.

CRINGOLI, G., QUESADA, A., and PAPPARELLA, V. 1991. Caryospora sp. (Apicomplexa: Eimeriidae) in common kestrel (Falco tinnunculus) Falconiformes: Falconidae. Rivista di Parassitologia 8: 39-46.

CRINGOLI, G., PAPPARELLA, V., QUESADA, A., and SCEBBA, S. 1989. Preliminary findings about the diffusion of Isospora and Caryospora species in wild birds. In: Proceedings of the 5th International Coccidiosis Conference, Tours (France), 17-20 October, 1989.

FREY, H., and KUTZER, E. 1982. Zur Diagnostik heimischer Greifvogelund Eulenparasiten. Praktische Tierarztliche 10: 894-902.

GOTTSCHALK, C. 1972. Beitrag zur Faunistik der Vogelkokzidien Thuringens und Sachsens. Beitraege zur Vogelkunde, Leipzig, 18: 61-69.

HELMBOLDT, C.F. 1967. An unidentified protozoan parasite in the kidney of the great-horned owl (Bubo virginianus). Bulletin of the Wildlife Diseases Association 3: 23-25.

HOARE, C.A. 1934. On the systematic status of Eumonospora tremula Allen, 1933 (Coccidiida Eimeriidae). Transactions of the American Microscopical Society 53: 6-7.

KLUH, P.N. 1994. Untersuchungen zur Therapie und Prophylaxe der Caryospora-Infektionen der Falken (Falconiformes: FAlconidae) mit Toltrazuril sowie die Beschreibung von zwei neuen Caryospora-Arten (C. megafalconis n. sp. und C. boeri n. sp.). PhD dissertation, Tierarztlichen Hochschule Hannover. 96 pp.

KUTZER, E. von, FREY, H., and KOTREMBA, J. 1980. Zur Parasitenfauna osterreichischer Greifvogel (Falconiformes). Angewandte Parasitologie 21: 183-205.

LINDSAY, D.S. and BLAGBURN, B.L. 1986. Caryospora uptoni n. sp. (Apicomplexa: Eimeriidae) from red-tailed hawks (Buteo jamaicensis borealis). Journal of Parasitology 72: 762-765.

LINDSAY, D.S. and BLAGBURN, B.L. 1989. Caryospora uptoni and Frenkelia sp.-like coccidial infections in red-tailed hawks, Buteo borealis. Journal of Wildlife Diseases 25: 407-409.

PELLERDY, L. 1967. Three new coccidia parasitic in Cuban birds (Protozoa: Sporozoa). Acta Zoologica Acadamiae Scientiarum Hungaricae 13: 227-230.

POELMA, F.G. and STRIK, W.J. 1966. Caryospora undata, Schwalbach 1959, a coccidium from the intestine of a guillemot, Uria aalge aalge, Pontoppidan, 1763). Tijdschrift voor Diergeneeskunde 91: 917-919.

SCHAFFRATH-BOER, A. 1983. Untersuchungen uber den endogenen Entwicklungszyklus von Caryospora kutzeri (Boer, 1982). Dissertation, Institute fur Parasitologie der Tierarztlichen Hochschule Hannover. 47 pp.

SCHELLNER, H.-P., and RODLER, S. 1971. Ein Trematoden-und Kokzidienbefall bei einem Luggerfalken (Falco juggar). Jahrbucher Deutscher Falkenorden 1970/1971: 90-93.

SCHWALBACH, G. 1959. Untersuchungen und Beobachtungen an Coccidien der Gattungen Eimeria, Isospora und Caryospora bei Vogeln mit einer Beschribung von sechzehn neuen Arten. Archiv fur Protistenkunde 104: 431-491.

STOCKDALE, P.H.G., and CAWTHORN, R.J. 1981. The coccidian Caryospora bubonis in the great horned owl (Bubo virginianus). Journal of Protozoology 28: 255-257.

SVOBODOVA, M. 1990. A preliminry study of monoxenous coccidia in passerine birds. Proceedings of the 20th annual conference, Protozoology section, Czechoslovakian Zoological Society. Acta Societatis Zoologicae Bohemoslovacae 56: 77.

SVOBODOVA, M. 1994. Isospora, Caryospora and Eimeria (Apicomplexa: Eimeriidae) in passeriform birds from Czech Republic. Acta Protozoologica 33: 101-108.

UPTON,S.J., CURRENT, W.L., and BARNARD, S.M. 1986. A review of the genus Caryospora Leger, 1904 (Apicomplexa: Eimeriidae). Systematic Parasitolology 8: 3-21.

UPTON, S.J., CAMPBELL, T.W., WEIGEL, M., and McKOWN, R.D. 1990. The Eimeriidae (Apicomplexa) of raptors: Review of the literature and description of new species of the genera Caryospora and Eimeria. Canadian Journal of Zoology 68: 1256-1265.

UPTON, S.J., ODELL, D.K., and WALSH, M.T. 1992. Description of the oocysts of Caryospora undata (Apicomplexa) from tufted puffins, Lunda cirrhata (Charadriiformes). Texas Journal of Science 44: 79-82.

UPTON, S.J. and SUNDERMANN, C.A. 1990. Caryospora: Biology. Chapter 10. In, Coccidiosis of Man and Domestic Animals, Long, P.L., editor. CRC Press, Boca Raton. pp. 187-204.

VARGHESE, T. and YAYABU, R. 1981. A survey of coccidian and helminth parasites of birds in Papua New Guinea with special reference to the Birds of Paradise. In, Wildlife Diseases of the Pacific Basin and other Countries, Proceedings of the 4th International Conference of the Wildlife Disease Association, August 25-28, 1981, Sydney, Australia, Fowler, M.E., editor. Wildlife Disease Association, Sydney. pp. 102-107.

VOLCAN, G.S., and MEDRANO, C.E. 1984. Esporozoario de aves Falconiformes en el estado Bolivar, Venezuela Caryospora arcayae n. sp. (Apicomplexa: Eimeriidae). In Cuadernos de Geografia Medica de Guayana. Edited by G.S. Volcan, C.V.G. Electrificacion del Caroni C.A./EDELCA, Venezuela, 1: 1-14.

VOLF, J. and MODRY, D. 1998. Prevalence of coccidia of genus Caryospora in captive birds of prey in the Czech republic. Bulletin of the Scandinavian Society of Parasitology 8: 86-87.

VOLF, J., KOUDELA, B., and MODRY, D. 2000. Two new species of Caryospora Leger, 1904 (Apicomplexa, Eimeriidae) from accipitrid raptors. Systematic Parasitology 46: 23-27.

WETZEL, R. and ENIGK, K. 1937. Caryospora falconis n. sp. (Eimeriidea) aus dem Wanderfalken. Sitzungsberichte Gesellschaft Naturforschender Freunde zu Berlin, Jan. 19, pp. 6-9.

WETZEL, R. and ENIGK, K. 1939. Beobachtungen uber parasitare Erkrankungen der Falken und Vorschlage zu ihrer Bekampfung. Gonderdrud aus der Zeitschrift Deutscher Falkenorden 1: 24-33.

YAKIMOFF, W.L. and MATIKASCHWILI, I.L. 1932. La coccidie du hibou. Bulletin de la Societe de Pathologie Exotique 25: 1045-1046.

YAKIMOFF, W.L. and MATSCHOULSKY, S.N. 1936. On the coccidia of birds of prey. Journal of the Royal Microscopical Society 56: 372-375.

YAKIMOFF, W.L. and MATSCHOULSKY, S.N. 1940. Coccidia in animals of the Zoological Gardens in Tashkent. Ecologischeskii Institut Parazitologischessi Sbornik, Akademii Nauk SSR, Leningrad 13: 236-248.

Class: Mammalia

Only a single species of mammalian caryosporan has been reported, and this species appears to be a pseudoparasite due to abnormal sporulation of isosporan oocysts. However, multiple reports have suggested that some members of the genus Caryospora may develop in ectopic sites in mammals, especially in the dermis (Douglas et al. 1991; Dubey and Speer 1991; Dubey et al. 1990; Sundermann et al. 1988, 1990, 1991). In the majority of naturally occurring cases, however, the exact organism cannot be identified (see Arcay 1982; Cable and Conaway 1953; Hrudka et al. 1983; Marcone 1908; Sangster et al. 1985; Shelton et al. 1968).

  1. Caryospora microti Saxe, Levine, and Ivens, 1960
    1. Host: Microtus pennsylvanicus (Rodentia)
    2. References: Saxe 1952; Saxe et al. 1960
    3. Remarks: This coccidian is the result of abnormal sporulation of an isosporan pseudoparasite. It should not be considered a valid species.

References for Caryospora spp. of Mammals

Arcay, L. 1982. Genital coccidiosis in the golden hampster (Cricetus cricetus), in Molecular and Biochemical Parasitology. Parasites- their World and ours, Muller, M., Gutteridge, W., and Kohler, P., Eds., Proceedings of the 5th International Congress of Parasitology (Toronto), August 7-14, Elsevier Biomedical Press, Toronto, pp. 404.

Cable, R.M. and Conaway, C.H. 1953. Coccidiosis of mammary tissue in the water shrew, Sorex palustris navigator. Journal of Parasitology 39: (suppl.) 30.

Douglas, R.J., Sundermann, C.A., Lindsay, D.S., and Mulvaney, D.R. 1991b. Development of Caryospora bigenetica (Apicomplexa: Eimeriidae) in porcine tissues. Journal of Protozoology 38: (Abstract 10): 2A.

Dubey, J.P. and Speer, C.A. 1991. Sarcocystis canis n. sp. (Apicomplexa: Sarcocystidae), the etiologic agent of generalized coccidiosis in dogs. Journal of Parasitology 77: 522-527.

Dubey, J.P., Black, S.S., Sangster, L.T., Lindsay, D.S., Sundermann, C.A., and Topper, M.J. 1990. Caryospora-associated dermatitis in dogs. Journal of Parasitology 76: 522-556.

Hrudka, F., Cawthorn, R.J., and Haigh, J.C. 1983. The occurence of coccidia (Eimeriidae) in epididymal semen of a wapiti (Cervus canadensis nelsoni). Canadian Journal of Zoology 61: 1693-1699.

Marcone, G. 1908. Sporozoen-Dermatosen des hundes. Zeitschrift fur Infektionskrankheiten, Parasitharekrankheiten und Hygiene der Haustiere 4: 5-32.

Sangster, L.T., Styler, E.L., and Hall, G.A. 1985. Coccidia associated with cutaneous nodules in a dog. Veterinary Pathology 22: 186-188.

Saxe, L.H. 1952. Observations on coccidia from Microtus pennsylvanicus. Journal of Protozoology 3 (Abstract):13.

Saxe, L.H., Levine, N.D., and Ivens, V. 1960. New species of coccidia from the meadow mouse, Microtus pennsylvanicus. Journal of Protozoology 7: 61-63.

Shelton, G.C., Kintner, L.D., and MacKintosh, D.O. 1968. A coccidia- like organism associated with subcutaneous granulomata in a dog. Journal of the American Veterinary Medical Association 152: 263-267.

Sundermann, C.A., Blagburn, B.L., Swango, L.J., and Boosinger, T. 1988. Dermal coccidiosis in dogs. Journal of the Alabama Academy of Sciences 59 (Abstract): 3.

Sundermann, C.A., Blagburn, B.L., Swango, L.J., and Boosinger, T.R. 1990. Development of Caryospora bigenetica (Apicomplexa) in canines. Journal of Protozoology 37(Supplement 27): 5A.

Sundermann, C.A., Lindsay, D.S., and Blagburn, B.L. 1991. Use of sulfadiazine to prevent dermal coccidiosis. Proceedings of the 44th annual meeting, Society of Protozoologists, Montana State University, Bozeman. 28 June - 2 July, 1991. Abstract 55, p. 41.

Class: Reptilia, Order: Chelonia

  1. Caryospora cheloniae Leibovitz, Rebell, and Boucher. 1978
    1. Host: Chelonia mydas (green sea turtle)
    2. References: Gordon et al. 1993; Laukner 1985; Leibovitz et al. 1978; Rebell et al. 1974; Sindermann 1988

References for the Caryospora spp. of Chelonia

Gordon, A.N., Kelly, W.R., and Lester, R.J. 1993. Epizootic mortality of free-living green turtles, Chelonia mydas, due to coccidiosis. Journal of Wildlife Diseases 29: 490-494.

Lauckner, G. 1985. Chapter 2. Diseases of Reptilia. In: Diseases of Marine Animals, Kinne, O., Editor. Volume 4, Part 2. Introduction, Reptilia, Aves, Mammalia. Biologische Anstalt Helgoland, Hamburg. pp. 552-626.

Leibovitz, L., Rebell, G., and Boucher, G.C. 1978. Caryospora cheloniae sp. n.: a coccidial pathogen of mariculture-reared green sea turtles (Chelonia mydas mydas). Journal of Wildlife Diseases 14: 269-275.

Rebell, G., Rywlin, A., and Ulrich, G. 1974. Coccidiosis in the green turtle (Chelonia mydas) in mariculture. In, Proceedings of the 5th Annual Meeting of the World Mariculture Association, Charleston, South Carolina, January 21-25, 1974. Avault, L.W., Jr., ed. Division of Continuing Education, Louisiana State University, Baton Rouge, 197-204.

Sindermann, C.J. 1988. Coccidian disease of green turtles. In, Disease Diagnosis and Control in North American Marine Aquaculture, Sindermann, C.J. and Lightner, D.V., editors. Elsevier Science Publishers, Amsterdam, pp. 384-385.

Class: Reptilia, Order: Sauria

  1. Caryospora ernsti Upton, Current, and Barnard, 1984
    1. Host: Anolis carolinensis (Green anole - Sauria)
    2. References: McAllister et al. 1994; Upton et al. 1984, 1986

  2. Caryospora gekkonis Chakravarty and Kar, 1947
    1. Host: Gekko gecko (Tokay - Sauria)
    2. Reference: Chakravarty and Kar 1947

  3. Caryospora varani Kaur and Oberoi, 1987 nomen dubia
    1. Host: The authors term the host "Varanus monitor" (Sauria); however, this is not a valid species. There are only four species of the genus Varanus in India, and the animal most likely represents either Varanus bengalensis or Varanus salvator.
    2. Reference: Kaur and Oberoi 1987
    3. Remarks: Based on oocyst size and the photomicrograph, this is clearly a misidentification and should be considered a nomen dubia

References for the Caryospora spp. of Sauria

Chakravarty, M. and Kar, A.B. 1947. Observations on two reptilian coccidia. Journal of the Royal Asiatic Society of Bengal, Science 12: 3-5.

Kaur, D. and Oberoi, R.K. 1987. Studies on protozoan gut fauna of Varanus monitor (carnivorous lizard). The Indian Zoologist 11: 35-39.

McAllister, C.T., Upton, S.J., and Trauth, S.E. 1994. New host and distribution records for coccidia (Apicomplexa: Eimeriidae) from North American lizards (Reptilia: Sauria). Journal of the Helminthological Society of Washington 61: 221-224.

Upton, S.J., Current, W.L., and Barnard, S.M., 1984b. A new species of Caryospora (Apicomplexa: Eimeriorina) from the green lizard, Anolis carolinensis. Transactions of the American Microscopical Society 103: 245-248.

Upton, S.J., Current, W.L., and Barnard, S.M. 1986. A review of the genus Caryospora Leger, 1904 (Apicomplexa: Eimeriidae). Systematic Parasitology 8: 321.

Class: Reptilia, Order: Serpentes

  1. Caryospora ahaetullae Modry and Koudela, 1994
    1. Host: Ahaetulla nasuta (Long nosed vine snake), Ahaetulla prasina (Green vine snake) (both Colubridae)
    2. References: Modry 1998; Modry and Koudela 1994, 1998

  2. Caryospora barnardae Upton, Freed, Burdick, and McAllister, 1990
    1. Host: Liopholidophis stumpffi (a Malagasy colubrid - Colubridae)
    2. Reference: Upton et al. 1990

  3. Caryospora bengalensis Mandal, 1976
    1. Host: Enhydris enhydris (Rainbow water snake - Colubridae)
    2. Reference: Mandal 1976

  4. Caryospora bigenetica Wacha and Christiensen, 1982
    1. Hosts: Agkistrodon contortrix (American copperhead), Crotalus adamanteus (Eastern diamondback rattlesnake), Crotalus atrox (Western diamondback rattlesnake), Crotalus horridus (Timber rattlensnake/Canebrake), Sistrurus catenatus (Massasauga), Sistrurus miliaris (Pigmy rattlesnake) (all Viperidae)
    2. References: Douglas et al. 1988, 1991a, 1991b; Dubey and Speer 1991; Dubey et al. 1990, 1991; Fayer and Dubey 1987; Lindsay and Dubey 1989; Lindsay et al. 1988; Modry 1998; Stibbs and Ongerth 1986; Sundermann 1990, 1991; Sundermann and Lindsay 1989a, 1989b, 1989c; Sundermann et al. 1988a, 1988b, 1988c, 1989, 1990, 1991; Tibbs et al. 1988; Upton and Barnard 1988; Upton and Sundermann 1990; Vitovec et al. 1997; Wacha and Christiensen 1981, 1982a, 1982b

  5. Caryospora brasiliensis Carini, 1932
    1. Hosts: Philodryas aestivus, Philodryas olfersi, Philodryas nattareri, Leimadophis poecilogyrus (all Colubridae)
    2. References: Carini 1932; Lainson and Shaw 1973

  6. Caryospora brygooi Upton, Freed, Burdick, and McAllister, 1990
    1. Host: Madagascarophis colubrinus (Madagascar cat-eye snake - Colubridae)
    2. Reference: Upton et al. 1990

  7. Caryospora carajasensis Lainson, Paiva do Nascimento, and Shaw, 1991
    1. Host: Oxyrhopus petola digitalis (Red-banded snake - Colubridae)
    2. Reference: Lainson et al. 1991

  8. Caryospora cobrae Nandi, 1985
    1. Host: Naja naja (Indian cobra - Elapidae)
    2. Reference: Nandi 1985

  9. Caryospora colubris Matuschka, 1984
    1. Host: Coluber hippocrepis (Horseshoe racer), Coluber jugularis (Whip snake), Coluber viridiflavus (Western whip snake); perhaps Eirenis decemlineata but this requires futher study (all Colubridae)
    2. References: Matuschka 1984b; Modry 1998; Paperna 1991; Paperna and Finkelman 1991
    3. Remarks: Experimental transmission attempts into SCID mice by Modry (1998) were unsuccessful.

  10. Caryospora constanciae Lainson, Paiva do Nascimento, and Shaw, 1991
    1. Host: Micrurus spixii spixii (Amazonian coral snake - Elapidae)
    2. Reference: Lainson et al. 1991

  11. Caryospora corallae Matuschka 1984
    1. Host: Corallus caninus (Emerald tree boa - Boidae)
    2. Reference: Matuschka 1984a

  12. Caryospora demansiae Cannon 1967
    1. Host: Demansia psammophis (Spotted-headed snake - Elapidae or Hydrophiidae, depending upon author)
    2. References: Cannon 1967; Rzepczyk 1976

  13. Caryospora dendrelaphis Cannon and Rzepczyk, 1974
    1. Host: Dendrelaphis punctulatus (Australian tree snake - Colubridae)
    2. References: Cannon and Rzepczyk 1974; Rzepczyk 1976

  14. Caryospora duszynskii Upton, Current, and Barnard, 1984
    1. Hosts: Elaphe guttata (Corn snake), Elaphe obsoleta (Rat snake), Lampropeltis calligaster (Prairie kingsnake) (all Colubridae)
    2. References: Lindsay et al. 1988; McAllister 1989; Modry 1998; Upton and McAllister 1990; Upton et al. 1984a
    3. Remarks: Experimental transmission attempts into SCID mice by Modry (1998) were unsuccessful.

  15. Caryospora epicratesi Lainson, Paiva do Nascimento, and Shaw, 1991
    1. Host: Epicrates cenchria cenchria (Brazilian rainbow boa - Boidae)
    2. References: Frank 1985; Lainson et al. 1991; Modry 1998
    3. Remarks: Experimental transmission attempts into SCID mice by Modry (1998) were unsuccessful.

  16. Caryospora gracilis Upton, McAllister, Trauth, and Bibb, 1992
    1. Host: Tantilla gracilis (Flatheaded snake - Colubridae)
    2. Reference: Upton et al. 1992b

  17. Caryospora heterodermus Upton, Freed, and Freed, 1992
    1. Host: Philothamnus heterodermus (Bush snake - Colubridae)
    2. Reference: Upton et al. 1992a

  18. Caryospora japonicum Matubayasi, 1936
    1. Host: Rhabdophis tigrinus (syn. Natrix tigrina) (Yamakagashi - Colubridae)
    2. References: Matubayasi 1936, 1937

  19. Caryospora jararacae Carini, 1939
    1. Host: Bothrops jararaca (Jararaca - Viperidae)
    2. References: Carini 1939; Lainson et al. 1991

  20. Caryospora kalimantanensis Modry and Koudela, 1997
    1. Host: Boiga dendrophila (Mangrove snake - Colubridae)
    2. References: Modry 1998; Modry and Koudela 1997

  21. Caryospora lampropeltis Anderson, Duszynski, and Marquardt, 1968
    1. Host: Lampropeltis calligaster (Prairie kingsnake - Colubridae)
    2. Reference: Anderson et al. 1968

  22. Caryospora legeri Hoare, 1933
    1. Synonym: Caryospora hermae Bray, 1960
    2. Synonym: Caryospora psammophi Bray, 1960
    3. Host: Psammophis sibilans (Hissing or Olive sand snake - Colubridae)
    4. References: Bray 1960; Daszak and Ball 2001; Hoare 1933

  23. Caryospora maculatus Upton, Freed, and Freed, 1992
    1. Host: Causus maculatus (Spotted night adder - Viperidae)
    2. Reference: Upton et al. 1992a

  24. Caryospora madagascariensis Upton, Freed, Burdick, and McAllister, 1990
    1. Hosts: Madagascarophis colubrinus (Madagascar cat-eye snake), Mimophis mahfalensis (a Malagasy colubrid) (both Colubridae)
    2. Reference: Upton et al. 1990

  25. Caryospora masticophis Upton, McAllister, and Trauth, 1994
    1. Hosts: Coluber constrictor priapus (Southern black racer - Colubridae); Masticophis flagellum flagellum (Eastern coachwhip - Colubridae)
    2. Reference: Upton et al. 1994

  26. Caryospora matatu Modry, Slapeta, and Koudela, 2002
    1. Host: Atheris ceratophorus (Horned bush-viper - Viperidae)
    2. Reference: Modry et al. 2002

  27. Caryospora maxima Modry, Koudela, Al-Oran, and Amr, 1999
    1. Host: Psammophis schokari (Forsskal's sand snake - Colubridae)
    2. References: Modry 1998; Modry et al. 1999

  28. Caryospora micruri Lainson, Paiva do Nascimento, and Shaw, 1991
    1. Host: Micrurus spixii spixii (Amazonian coral snake - Elapidae)
    2. Reference: Lainson et al. 1991

  29. Caryospora minuta Upton, Freed, Burdick, and McAllister, 1990
    1. Host: Lioheterodon madagascariensis (Madagascar hognose snake - Colubridae)
    2. References: Modry 1998; Upton et al. 1990
    3. Remarks: Experimental transmission attempts into SCID mice by Modry (1998) were unsuccessful.

  30. Caryospora najadae Matuschka 1986
    1. Host: Coluber najadum (Dahl's whip snake - Colubridae)
    2. Reference: Matuschka 1986a

  31. Caryospora najae Matuschka 1982
    1. Host: Naja pallida (syn. Naja nigricollis pallida) (Red spitting cobra - Elapidae)
    2. Reference: Matuschka 1982

  32. Caryospora paraensis Lainson, Paiva do Nascimento, and Shaw, 1991
    1. Host: Oxyrhopus petola digitalis (Colubridae)
    2. Reference: Lainson et al. 1991

  33. Caryospora peruensis Upton, Freed, and Freed, 1989
    1. Host: Oxybelis argenteus (Argentine vine snake - Colubridae)
    2. Reference: Upton et al. 1989

  34. Caryospora pseustesi Lainson, Paiva do Nascimento, and Shaw, 1991
    1. Host: Pseustes sulphureus sulphureus (Colubridae)
    2. Reference: Lainson et al. 1991

  35. Caryospora regentensis Daszak and Ball, 2001
    1. Host: Dendroaspis viridis (Green mamba - Elapidae)
    2. Reference: Daszak and Ball 2001

  36. Caryospora relictae Telford, 1997
    1. Host: Tantilla relicta (Peninsular crowned snake - Colubridae)
    2. Reference: Telford 1997

  37. Caryospora serpentis Upton, Freed, Burdick, and McAllister, 1990
    1. Hosts: Lioheterodon madagascariensis (Madagascar hognose snake), Madagascarophis colubrinus (Madagascar cat-eye snake), Mimophis mahfalensis (a Madagascar colubrid) (all Colubridae)
    2. References: Modry 1998; Upton et al. 1990
    3. Remarks: Experimental transmission attempts into SCID mice by Modry (1998) were unsuccessful.

  38. Caryospora simplex Leger, 1904
    1. Synonym: Karyospora simplex Leger, 1904, lapsus
    2. Hosts: Vipera ammodytes (Long-nosed viper), Vipera aspis (Asp viper), Vipera berus (European viper), Vipera kaznakovi (Kaznakov's viper), Vipera palestinae (Palestine viper), Vipera russelli (Russell's viper), Vipera xanthina (Palestine viper) (all Viperidae)
    3. References: Augustine and Danforth 1985; Barnard 1986; Lavier 1939; Leger 1904, 1911; Matuschka 1986b; Modry 1998; Modry et al. 1997; Upton 1983; Upton and Barnard 1986; Upton et al. 1983a, 1983b, 1983c, 1984c, 1984d, 1984e, 1984f, 1985; Vitovec et al. 1997; Wilber et al. 1995
    4. Remarks: Cross transmission studies by Modry (1998) resulted in successful transmissions to Cerastes cerastes (Horned viper), Echis coloratus (Arabian viper), and Pseudocerastes persicus (Palestine or Persian horned viper).

  39. Caryospora sp. of Hoge, 1991
    1. Host: Bothrops jararacussu (Jararacussu - Viperidae)
    2. Reference: Hoge 1991

  40. Caryospora sp. of Hoge, 1991
    1. Host: Bothrops neuwiedi (Jararaca pintada or Wied's lance-head - Viperidae)
    2. Reference: Hoge 1991

  41. Caryospora sp. of Hoge, 1991
    1. Host: Tomodon dorsatus (Cobra-espada or Jararaca-da-chuva - Colubridae)
    2. Reference: Hoge 1991

  42. Caryospora sp. of Modry, 1998
    1. Host: Coluber ventromaculata (Colubridae)
    2. Reference: Modry 1998

  43. Caryospora sp. 1 of Modry, 1998
    1. Host: Atheris nitschei (Nitsche's bush viper - Viperidae)
    2. Reference: Modry 1998
    3. Remarks: Experimentally inoculated SCID mice became infected but failed to transmit the infection to Vipera ammodytes (Long-nosed viper).

  44. Caryospora sp. 2 of Modry, 1998
    1. Host: Calloselasma rhodostoma (Malayan pit viper - Viperidae)
    2. Reference: Modry 1998
    3. Remarks: Experimentally inoculated SCID mice became infected but failed to transmit the infection to Vipera ammodytes (Long-nosed viper), Crotalus atrox (Western diamondback rattlesnake), or Gloydius blomhoffi (syn. Agkistrodon blomhoffi) (Mamushi)

  45. Caryospora sp. of Upton and Sundermann, 1990
    1. Host: Bothrops lateralis (Yellow-lined pit viper - Viperidae)
    2. Reference: Upton and Sundermann 1990

  46. Caryospora spp. of Hoge, 1991
    1. Hosts: Crotalus durissus ssp. (a new world rattlesnake), Crotalus durissus terrificus (South American rattlesnake) Viperidae)
    2. Reference: Hoge 1991
    3. Remarks: Seven different morphologic types were measured from a total of 5 different specimens of Crotalus durissus. The differences between the isolates are relatively minor and they may represent only one or two species.

  47. Caryospora tantillae Telford, 1997
    1. Host: Tantilla relicta (Peninsular crowned snake - Colubridae)
    2. Reference: Telford 1997

  48. Caryospora telescopis Matuschka 1986
    1. Host: Telescopus fallax (Cat snake - Colubridae)
    2. Reference: Matuschka 1986b

  49. Caryospora veselyi Modry and Koudela, 1998
    1. Host: Ahaetulla nasuta (Long nosed vine snake - Colubridae)
    2. References: Modry 1998; Modry and Koudela 1998

  50. Caryospora weyerae Bray, 1960
    1. Host: Psammophis sibilans (Hissing or Olive sand snake - Colubridae)
    2. Reference: Bray 1960

  51. Caryospora zuckermanae Bray, 1960
    1. Host: Coluber ravergieri (Colubridae)
    2. Reference: Bray 1960

References for Caryospora spp. of Serpentes

Anderson, D.R., Duszynski, D.W., and Marquardt, W.C. 1968. Three new coccidia (Protozoa: Telosporea) from kingsnakes, Lampropeltis spp., in Illinois, with a redescription of Eimeria zamenis Phisalix, 1921. Journal of Parasitology 54: 577-581.

Augustine, P.C. and Danforth, H.D. 1985. Hybridoma antibodies reveal differences in antigenic composition of coccidia refractile bodies. Abstract from the 60th annual meeting of the American Society of Parasitologists, August 4-8, Athens, Georgia, p. 56.

Barnard, S.M. 1986. Color atlas of reptilian parasites. Part I. Protozoans. Compendium on Continuing Education for the Practicing Veterinarian 8: 145-150.

Bray, R.S. 1960. On the parasitic protozoa of Liberia. III. Three new species of Caryospora Leger, 1904 from the hissing sand snake Psammophis sibilans phillipsi and a note on a Caryospora in Coluber ravergieri nummifer from Israel. Journal of Protozoology 7: 314-320.

Cannon, L.R.G. 1967. Caryospora demansiae sp. nov. (Sporozoa: Eimeriidae) from an Australian snake Demansia psammophis (Elapidae). Parasitology 57: 221-226.

Cannon, L.R.G. and Rzepczyk, C.M. 1974. A new species of Caryospora Leger, 1904 (Sporozoa: Eimeriidae) from an Australian snake. Parasitology 69: 197-200.

Carini, A. 1932. Caryospora brasiliensis n. sp., parasita do intestino da cobra verde, Chlorosoma aestivum. Revista de Biologia e Hygiene 3: 85-87.

Carini, A., Sobre uma cariospora da Jararaca. Archivos de Biologia, Sao Paulo 23: 41-42.

Daszak, P. and Ball, S.J. 2001. A description of two new species of coccidia (Apicomplexa: Eimeriidae) from African reptiles with nomenclatural corrections for two Caryospora and one Eimeria species from snakes. Folia Parasitologica (Praha) 48: 1-6.

Dollfus, R.-Ph. 1961. Station experimentale de parasitologie den Richelieu (Indre-et-Loire) contribution a la faune parasitaire reginale. Annales de Parasitologie Humaine et Comparee 36: 171-355.

Douglas, R.J., Sundermann, C.A., and Lindsay, D.S. 1988. New modes of transmission of Caryospora bigenetica (Apicomplexa: Eimeriidae). Journal of the Alabama Academy of Sciences 59: 128.

Douglas, R.J., Sundermann, C.A., and Lindsay, D.S. 1991a. Effects of route of inoculation on the site of development of Caryospora bigenetica (Apicomplexa: Eimeriidae). Journal of Parasitology 77: 755-757.

Douglas, R.J., Sundermann, C.A., Lindsay, D.S., and Mulvaney, D.R. 1991b. Development of Caryospora bigenetica (Apicomplexa: Eimeriidae) in porcine tissues. Journal of Protozoology 38:(Abstract 10): 2A.

Dubey, J.P. and Speer, C.A. 1991. Sarcocystis canis n. sp. (Apicomplexa: Sarcocystidae), the etiologic agent of generalized coccidiosis in dogs. Journal of Parasitology 77: 522-527.

Dubey, J.P., Black, S.S., Sangster, L.T., Lindsay, D.S., Sundermann, C.A., and Topper, M.J. 1990. Caryospora-associated dermatitis in dogs. Journal of Parasitology 76: 522-556.

Dubey, J.P., Davis, S.W., Speer, C.A., Bowmann, D.D., de Lahunta, A., Grandstrom, D.E., Topper, M.J., Hamis, A.N., Cummings, J.F., and Suter, M.M. 1991. Sarcocystis neurona n. sp. (Protozoa: Apicomplexa), the etiologic agent of equine protozoal myeloencephalitus. Journal of Parasitology 77: 212-218.

Fayer, R. and Dubey, J.P.. 1987. Comparative epidemiology of coccidia: clues to the etiology of equine protozoal myeloencephalitis. International Journal for Parasitology 17: 615-620.

Frank, W. 1985. Amphibien und Reptilian. In, Heimtierkrankheiten. Isenbugel, E. and Frank, W. eds. Verlag Eugene Ulmer, Stuttgart. pp. 162-382.

Hoare, C.A. 1933. Studies on some new ophidian and avian coccidia from Uganda, with a revision of the classification of the Eimeriidoa. Parasitology 25: 359-388.

Hoge, A.Y.A. 1991. Ocorrencia de coccidias intestinais (Subordem Eimeriorina) em algumas serpentes Brasileiras (Colubridae e Crotalinae). PhD Dissertation, Universidade de Sao Paulo. 55 pp.

Lainson, R. and Shaw, J.J. 1973. Coccidia of Brazilian snakes: Isospora decipiens, Eimeria micruri, E. liophi and E. leimadophi spp. n., with redescriptions of Caryospora brasiliensis Carini, 1932 and Eimeria poecilogyri Carini, 1933. Journal of Protozoology 20: 358-362.

Lainson, R., Paiva do Nascimento, F., and Shaw, J.J. 1991. Some new species of Caryospora (Apicomplexa: Eimeriidae) from Brazilian snakes, and a re-description of C. jararacae Carini, 1939. Memorias do Instituto Oswaldo Cruz 86: 349-364.

Lavier, G. 1939. Sur une coccidie atypique de Vipera aspis. Annales de Parasitologie Humaine et Comparee 17: 37-44.

Leger, L.1904. Protozoaires parasites des Viperes (Vipera aspis). Bulletin Mensuel, Association Francaise pour l'avancement des Sciences (Abstract) 9: 268.

Leger, L. 1911. Caryospora simplex, coccidie monosporee et la classification des coccidies. Archiv fur Protistenkunde 22: 71-88.

Lindsay, D.S. and Dubey, J.P. 1989. Immunohistochemical diagnosis of Neospora caninum in tissue sections. American Journal of Veterinary Research 50: 1981-1983.

Lindsay, D.S. and Sundermann, C.A., 1989. Recent advances in the biology of the coccidian genus Caryospora. Journal of Veterinary Parasitology 3: 1-5.

Lindsay, D.S., Sundermann, C.A., and Blagburn, B.L. 1988. Caryocyst-like host cell formation by Caryospora duszynskii (Apicomplexa: Eimeriidae) in human fetal lung cell cultures. Journal of Protozoology 35: 32-33.

Lindsay, D.S., Sundermann, C.A., Blagburn, B.L., Upton, S.J., and Barnard, S.M. 1988. Fatal Caryospora bigenetica (Apicomplexa: Eimeriidae) infections in cotton rats, Sigmodon hispidus. Journal of Parasitology 74: 838-841.

Mandal, R. 1976. Caryospora bengalensis sp. n. and Eimeria fibrilosa sp. n., new coccidia (Protozoa: Eucoccida) from a fresh-water snake, Enhydris enhydris (Schneider). Acta Protozoologica 15: 405-410.

Matubayasi, H. 1936. On a new coccidia, Koidzumiella natrix n. g., n. sp., parasitic in Japanese snakes; with some remarks on two other ophidian coccidia of the genus Caryospora and Isospora. Keio-Igaku 16: 1719-1729

Matubayasi, H. 1937. Studies on parasitic protozoa in Japan II. Coccidia parasitic in snakes; with special remarks on Tyzzeria (Koidzumiella) natrix, parasitic in Natrix tigrina. Annotationes Zoologicae Japonenses 16: 255-275.

Matuschka, F.-R. 1982. Caryospora najae sp. n. (Apicomplexa: Sporozoea, Eimeriidae) from the spitting cobra, Naja nigricollis pallida (Serpentes: Elapidae). Journal of Parasitology 68: 1149-1153.

Matuschka, F.-R. 1984a. Caryospora corallae n. sp. (Apicomplexa: Sporozoea, Eimeriidae) from the emerald tree boa Corallus caninus (Serpentes: Boidae). Journal of Protozoology 31: 358-359.

Matuschka, F.-R. 1984b. Description of Caryospora colubris sp. n. (Apicomplexa: Eimeriidae) from the western whip snake, Coluber viridiflavus (Serpentes: Colubridae). Canadian Journal of Zoology 62: 1525-1527.

Matuschka, F.-R. 1986a. Caryospora najadae sp. n. (Apicomplexa: Eimeriidae) from Dahl's whip snake, Coluber najadum (Serpentes: Colubridae). Zeitschrift fur Parasitenkunde 72: 181-183.

Matuschka, F.-R. 1986b. Caryospora telescopis sp. n. (Apicomplexa: Eimeriidae) from the cat snake, Telescopus fallax (Serpentes: Colubridae), and Caryospora spec. from the West Caucasian viper, Vipera kaznakovi (Serpentes: Viperidae). Archiv fur Protistenkunde 132: 93-99.

McAllister, C.T. 1989. Systematics of coccidian parasites (Apicomplexa) from amphibians and reptiles in Northcentral Texas. PhD dissertation, University of North Texas, Denton, Texas, 152 pp.

Modry, D. 1998. The biology and pathogenicity of caryosporan coccidia. PhD dissertation, University of Veterinary and Pharmaceutical Sciences Brno. 52 pp. plus appendices.

Modry, D., and Koudela, B. 1994. Caryospora ahaetullae sp. n. (Apicomplexa: Eimeriidae) from the long nosed vine snake, Ahaetulla nasuta (Serpentes: Colubridae). Folia Parasitologica (Praha) 41: 233-235.

Modry, D., and Koudela, B. 1997. Caryospora kalimantanensis sp. n. (Apicomplexa: Eimeriidae) from the mangrove snake, Boiga dendrophila (Serpentes: Boiginae) from Kalimantan. Folia Parasitologica (Praha) 44: 195-199.

Modry, D., and Koudela, B. 1998. Coccidia of the genus Caryospora (Apicomplexa: Eimeriidae) from the long nosed vine snake, Ahaetulla nasuta (Serpentes: Colubridae: Boiginae) from southern India, with a description of C. veselyi. Folia Parasitologica (Praha) 45: 167-169.

Modry, D., Koudela, B., Al-Oran, R.M., and Amr, Z.S. 1999. Caryospora maxima sp. n. (Apicomplexa: Eimeriidae), new coccidium from Forskal's sand snake, Psammophis schokari (Serpentes: Colubridae) from lava desert of Jordan. Folia Parasitologica (Praha) 46: 96-98.

Modry, D., Koudela, B., Volf, J., Necas, P., and Hudcovic, T. 1997. Vipera berus and V. ammodytes (Serpentes: Viperidae) represent new hosts for Caryospora simplex (Apicomplexa: Eimeriidae) in Europe. Folia Parasitologica (Praha) 44: 99-102.

Modry, D., Slapeta, J.R., and Koudela, B. 2002. Caryospora matatu n. sp., a new coccidian parasite (Apicomplexa: Eimeriidae) from the horned bush-viper, Atheris ceratophorus Wener, 1895 from Tanzania. Parasite 9: 341-344.

Nandi, R. Caryospora cobrae n. sp. from the Indian cobra Naja naja Linnaeus. Journal of Protozoology 32: 204-205.

Paperna, I. 1991. Electron microscopic studies of Caryospora colubris development in the Syrian black snake Coluber jugularis. Annales de Parasitologie humaine et Comparee 66: 139-143.

Paperna, I., and Finkelman, S. 1991. Development of Caryospora colubris (Apicomplexa: Eimeriidae) in the Syrian black snake Coluber jugularis L. Annales de Parasitologie humaine et Comparee 66: 91-95.

Pellerdy, L. 1974. Coccidia and Coccidiosis, Verlag Paul Parey, Berlin, 959 pp.

Rzepczyk, C.M. 1976. Studies on Sarcocystis with particular reference to the life history. Phd dissertation, University of Queensland. 247 pp.

Stibbs, H.H. and Ongerth, J.E. 1986. Immunofluorescence detection of Cryptosporidium oocysts in fecal smears. Journal of Clinical Microbiology 24: 517-521.

Sundermann, C.A. 1990. Long term cultivation of Caryosporabigenetica (Apicomplexa). Journal of the Alabama Academy of Sciences 61(Abstract): 138.

Sundermann, C.A. 1991. Continuous in vitro growth of a coccidian of snakes and mammals. Journal of Protozoology 38 (Abstract 86): 15A.

Sundermann, C.A., Blagburn, B.L., Swango, L.J., and Boosinger, T. 1988a. Dermal coccidiosis in dogs. Journal of the Alabama Academy of Sciences 59 (Abstract): 3.

Sundermann, C.A., Blagburn, B.L., Swango, L.J., and Boosinger, T.R. 1990. Development of Caryospora bigenetica (Apicomplexa) in canines. Journal of Protozoology 37(Supplement 27): 5A.

Sundermann, C.A. and Lindsay, D.S. 1989a. Complete development of Caryospora (Apicomplexa: Eimeriidae) in cultured mammalian cells. Journal of Protozoology 36 (Supplement 82): 14A.

Sundermann, C.A. and Lindsay, D.S. 1989b. Investigation of the caryocyst of Caryospora bigenetica (Apicomplexa: Eimeriidae). Journal of the Alabama Academy of Sciences 60 (Abstract): 130.

Sundermann, C.A. and Lindsay, D.S. 1989c. Ultrastructure of in vivo-produced caryocysts containing the coccidian Caryospora bigenetica (Apicomplexa: Eimeriidae). Journal of Protozoology 36: 81-86.

Sundermann, C.A., Lindsay, D.S., and Blagburn, B.L. 1988b. Experimental infections and ultrastructure of Caryospora bigenetica in cotton rats. Journal of the Alabama Academy of Sciences. 59 (Abstract): 129.

Sundermann, C.A., Lindsay, D.S., and Blagburn, B.L. 1991. Use of sulfadiazine to prevent dermal coccidiosis. Proceedings of the 44th annual meeting, Society of Protozoologists, Montana State University, Bozeman. 28 June - 2 July, 1991. Abstract 55, p. 41.

Sundermann, C.A., Lindsay, D.S., Blagburn, B.L., Bailey, M.A., and Wester, E.E. 1989. Serial transmission of Caryospora bigenetica Wacha and Christiansen, 1982 (Apicomplexa: Eimeriidae) between different species of rodents. Journal of Parasitology 75: 327-329.

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Upton, S.J. and Barnard, S.M. 1988. Development of Caryospora bigenetica (Apicomplexa: Eimeriorina) in experimentally infected mice. International Journal for Parasitology 18: 15-20.

Upton, S.J. and Barnard, S.M. 1986. Experimental transmission of Caryospora simplex (Apicomplexa: Eimeriidae) to Palestine vipers, Vipera xanthina palestinae (Serpentes: Viperidae). Journal of Protozoology 33: 129-130.

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Upton, S.J., Current, W.L., and Barnard, S.M., 1984a. A new species of Caryospora (Apicomplexa: Eimeriorina) from Elaphe spp. (Serpentes: Colubridae) of the southeastern and central United States. Transactions of the American Microscopical Society 103: 240-244.

Upton, S.J., Current, W.L., and Barnard, S.M. 1986. A review of the genus Caryospora Leger, 1904 (Apicomplexa: Eimeriidae). Systematic Parasitology 8: 321.

Upton, S.J., Current, W.L., Barnard, S.M., and Ernst, J.V. 1984c. In vitro excystation of Caryospora simplex (Apicomplexa: Eimeriorina). Journal of Protozoology 31: 293-297.

Upton, S.J., Current, W.L., Ernst, J.V., and Barnard, S.M.1983a. Caryospora simplex: Extraintestinal development in mice. Association of Southeastern Biologists Bulletin 30: 88.

Upton, S.J., Current, W.L., Ernst, J.V., and Barnard, S.M. 1983b.Development of the viperid parasite, Caryospora simplex in mice. Journal of the Alabama Academy of Sciences 54: 114.

Upton, S.J., Current, W.L., Ernst, J.V., and Barnard, S.M. 1984d. Extraintestinal development of Caryospora simplex (Apicomplexa: Eimeriidae) in experimentally infected mice, Mus musculus. Journal of Protozoology 31: 392-398.

Upton, S.J., Ernst, J.V., Barnard, S.M., and Current, W.L. 1983c. Redescription of the oocysts of Caryospora simplex (Apicomplexa: Eimeriidae) from an Ottoman viper, Vipera xanthina xanthina. Transactions of the American Microscopical Society 102: 258-263.

Upton, S.J., Freed, P.S., Burdick, D.A., and McAllister, C.T. 1990. Seven new species of coccidia (Apicomplexa: Eimeriorina) from reptiles in Madagascar. Canadian Journal of Zoology 68: 2368-2375.

Upton, S.J., Freed, P.S., and Freed, M.G. 1989. Two new species of coccidia (Apicomplexa: Eimeriorina) from Peruvian reptiles. Transactions of the American Microscopical Society 108: 96-100.

Upton, S.J., Freed, P.S., and Freed, M.G. 1992a. Two new species of Caryospora (Apicomplexa) from snakes in Cameroon, with redescriptions of two coccidians from Hemidactylus mabouia (Gekkonidae). Systematic Parasitology 23: 135-140.

Upton, S.J., Haynes, T.B., Current, W.L., and Barnard, S.M. 1984e. Development of Caryospora simplex (Apicomplexa: Eimeriidae) from sporozoites to oocysts in human embryonic lung cell culture. Journal of Protozoology 31: 398-402.

Upton, S.J., Haynes, T.B., Current, W.L., and Barnard, S.M. 1984f. Development of Caryospora simplex in human embryonic lung cell culture. Association of Southeastern Biologists Bulletin 31: 88.

Upton, S.J., McAllister, C.T., and Trauth, S.E. 1994g. Caryospora masticophis n. sp. (Apicomplexa) from Masticophis flagellum and Coluber contrictor (Serpentes) in Arkansas, U.S.A. Transactions of the American Microscopical Society 113: 395-399.

Upton, S.J., Lindsay, D.S., Current, W.L., and Barnard, S.M. 1985. Mouse-to-mouse transmission of Caryospora simplex (Apicomplexa: Eimeriidae). Journal of Parasitology 71: 395-396.

Upton, S.J., McAllister, C.T., Trauth, S.E., and Bibb, D.K. 1992b. Description of two new species of coccidia (Apicomplexa: Eimeriorina) from flat-headed snakes, Tantilla gracilis (Serpentes: Colubridae) and reclassification of misnomer species within the genera Isospora and Sarcocystis from snakes. Transactions of the American Microscopical Society 111: 50-60.

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